Understanding and modeling microbial responses and feedbacks to climate change is hampered by a lack of a framework in the pelagic environment by which to link local mechanism to large scale patterns. Where terrestrial ecology draws from landscape theory and practice to address issues of scale, the pelagic seascape concept is still in its infancy. We have applied the patch mosaic paradigm of landscape ecology to the study of the seasonal and interannual variability of the North Pacific to facilitate comparative analysis between pelagic ecosystems and provide spatiotemporal context for eulerian time-series studies. Using multivariate, 13-year climatologies of sea surface temperature, photosynthetically active radiation, and chlorophyll a derived from remote sensing observations, we classified hierarchical seascapes at monthly and interannual scales. These dynamic, objectively-determined seascapes offer improved hydrographic coherence relative to oceanic regions with subjectively defined and static boundaries (Chapter 2) and represent unique biogeochemical functioning (Chapter 2) and microbial communities (Chapter3). Furthermore they provide consilience between satellite studies and in situ observations (Chapter 4) and allow for objective comparison of ecosystem forcing (Chapters, 4 and 5).
In Chapter 2, we rigorously tested the assumption that satellite-derived seascapes describe regions of biogeochemical coherence. The seasonal cycle of the North Pacific was characterized at three levels of spatiotemporal hierarchy and broader relevance of monthly –resolved seascapes was assessed through analysis of variance (ANOVA) and multiple linear regression (MLR) analyses of nutrient, primary productivity, and pCO₂ data. Distinct nutrient and primary productivity regimes were well-characterized in the coarsest two levels of hierarchy (ANOVA, R² = 0.5-0.7). Finer scale partitioning was more relevant for pCO₂. MLR analyses revealed differential forcing on pCO₂ across seascapes and hierarchical levels and a 33 % reduction in mean model error with increased partitioning (from 18.5 µatm to 12.0 µatm pCO₂).
In Chapter 3 we verified the seascapes with in situ collections of microbial abundance and structure. Flow cytometry data was collected from two long term time series and several cruises spanning thousand kilometers of the NE Pacific; these data allowed us to quantify spatiotemporal patterns. In addition, multiple response permutation analysis revealed differences in community structure across discrete seascapes, in terms of both absolute and relative abundances. Principal component analysis of the assemblage supported seascape divisions and revealed structure along environmental gradients with strong associations with chlorophyll a and sea surface temperature and, to a lesser extent, with mixed layer depth and mean photosynthetically active radiation in the mixed layer. Differences of assemblage structure between seascapes and strength of environmental forcing were strong in the subarctic and transition zones, but less pronounced in the subtropics, suggesting satellite-detected changes in bulk properties that may be associated with local physiology or interannual shifts in seascape boundaries.
Based on the work presented in Chapter 4, we discovered that interannual shifts in the boundaries of a transition seascape and two distinct oligotrophic subtropical seascapes affect the variability observed at benchmark time series Station ALOHA; the latter two seascapes oscillate in their contributions to the expansion of the entire subtropics. On interannual scales, in situ phytoplankton abundance (as measured by chl-a), net primary productivity (NPP), and the relative abundance of eukaryotic phytoplankton and Synechococcus sp. increased during periods of encroachment by the transition seascape. Conversely, the relative abundance of Prochlorococcus increased and chl –a and NPP decreased when the highly oligotrophic seascape encroached on Station ALOHA. The dynamic range (~6 million km²) of subtropical expansion is born almost entirely by the transition zone - resulting in a transfer of ~1.2 Pg of total primary C production between a system primed for export production and one dominated by the microbial loop.
In Chapter 5, we investigated multiple factors that contribute to the effectiveness of the biological pump in the transition seascape. Near-continuous measurements of net primary production (NPP), net community production (NCP) and several ecophysiological variables were collected in across subarctic, transition, and subtropical seascapes of the Northeast Pacific during August and September of 2008. Mesoscale processes and shifts in community structure contributed to high export efficiency in the subtropical seascape; the convergence of assemblage structure, high biomass, moderate NPP: NCP and high NCP contributed to biologically mediated air-sea exchange in the transition seascape. Furthermore, NPP and NCP were strongly spatially coupled in both the transition (r[subscript 1, 39]=0.70; p<0.0001) and subtropical seascapes (r[subscript 1, 45]= 0.68, p<0.0001), suggesting the possibility for empirical modeling efforts.
This dissertation provides a first step to characterize the seascape variability in the NE Pacific and to understand the modulation of primary and export production in a critical transition region. The multivariate seascape approach described here provides spatiotemporal context for in situ studies and allows objective comparisons of systems' responses to climatic forcing. An integrated ocean observing system will require insight from in situ observations and experiments, ecosystem models, and satellite remote sensing. The results highlighted in this dissertation suggest that the pelagic seascape framework, through its capacity to scale both context and mechanism, may serve as an important and unifying component of such an observing system.